Human burials at the Kisese II rockshelter, Tanzania

Received: 31 August 2020 Revised: 4 December 2020 Accepted: 29 January 2021 DOI: 10.1002/ajpa.24253 RESEARCH ARTICLE Human burials at the Kisese II rockshelter, Tanzania | Elizabeth A. Sawchuk2,3 | Amandus Kwekason4 | Myra F. Laird1 Audax Z. P. Mabulla5 | Emmanuel Ndiema6 | Christian A. Tryon6,7 | Jason E. Lewis3,8 | Kathryn L. Ranhorn9 1 Department of Integrative Anatomical Sciences, University of Southern California, Los Angeles, California, USA 2 Department of Anthropology, University of Alberta, Edmonton, Alberta, Canada 3 Department of Anthropology, Stony Brook University, Stony Brook, New York, USA 4 National Museums of Tanzania, Dar es Salaam, Tanzania 5 Department of Archaeology and Heritage Studies, University of Dar es Salaam, Dar es Salaam, Tanzania 6 Department of Anthropology, University of Connecticut, Storrs, Connecticut, USA 7 Human Origins Program, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA 8 Turkana Basin Institute, Stony Brook University, Stony Brook, New York, USA 9 Institute of Human Origins, School of Human Evolution and Social Change, Arizona State University, Tempe, Arizona, USA Correspondence Myra F. Laird, Department of Integrative Anatomical Sciences, University of Southern California, 1333 San Pablo Street, Los Angeles, CA 90033, USA. Email: [email protected] Funding information Fulbright-Hays; Leakey Foundation; National Science Foundation, Grant/Award Number: DGE 0801634; Social Sciences and Humanities Research Council of Canada, Grant/Award Number: 767-2012-1903; NSF Archaeometry program grant to Kennett and Culleton, Grant/ Award Number: (BCS-1460369; Dental metric data were collected under NACOSTI, Grant/ Award Numbers: NACOSTI/P/14/1876/1410, NCST/5/002/R/576; Kenyan National Commission for Science, Technology, and Innovation (NACOSTI); Tanzania Commission for Science and Technology (COSTECH), Grant/Award Numbers: 2013-223-NA2014-101, 2015-120-NA-2015-24, 2015-116-ER-2015-212, 2015-115-ER2013-122, 2014-233-NA-2013-122 Abstract Objectives: The Late Pleistocene and early Holocene in eastern Africa are associated with complex evolutionary and demographic processes that contributed to the population variability observed in the region today. However, there are relatively few human skeletal remains from this time period. Here we describe six individuals from the Kisese II rockshelter in Tanzania that were excavated in 1956, present a radiocarbon date for one of the individuals, and compare craniodental morphological diversity among eastern African populations. Materials and Methods: This study used standard biometric analyses to assess the age, sex, and stature of the Kisese II individuals. Eastern African craniodental morphological variation was assessed using measures of dental size and a subset of Howells' cranial measurements for the Kisese II individuals as well as early Holocene, early pastoralist, Pastoral Neolithic, and modern African individuals. Results: Our results suggest a minimum of six individuals from the Kisese II collections with two adults and four juveniles. While the dating for most of the burials is uncertain, one individual is directly radiocarbon dated to
7.1 ka indicating that at least one burial is early Holocene in age. Craniodental metric comparisons indicate that the Kisese II individuals extend the amount of human morphological diversity among Holocene eastern Africans. Conclusions: Our findings contribute to a growing body of evidence that Late Pleistocene and early Holocene eastern Africans exhibited relatively high amounts of morphological diversity. However, the Kisese II individuals suggest morphological This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and distribution in any medium, provided the original work is properly cited, the use is non-commercial and no modifications or adaptations are made. © 2021 The Authors. American Journal of Physical Anthropology published by Wiley Periodicals LLC. Am J Phys Anthropol. 2021;1–14. wileyonlinelibrary.com/journal/ajpa 1 LAIRD ET AL. 2 similarity at localized sites potentially supporting increased regionalization during the early Holocene. KEYWORDS early Holocene, eastern Africa, morphological variation 1 | I N T RO DU CT I O N et al., 2019) and Pastoral Neolithic (PN) herders in southern Kenya and northern Tanzania
4–1.5 ka (Ambrose, 2001; Bower, 1991; Human (and more broadly, hominin) skeletal remains from eastern Sawchuk et al., 2018). Africa play a central role in discussions about human origins, the evo- Previous research on Late Pleistocene LSA individuals indicates lution of morphological diversity within Homo sapiens, and population at least some populations exhibit patterns of morphological diver- dispersals and interactions associated with the spread of animal and sity distinct from recent sub-Saharan Africans (Crevecoeur plant domesticates. Yet despite their importance, little is known about et al., 2009; Crevecoeur et al., 2016; Grine et al., 2007; Mounier human morphological variation from the region. This is because (1) there et al., 2018; Reiner et al., 2017; Stojanowski, 2014; Tryon are relatively few sites with human remains, (2) bones (when preserved) et al., 2015). This could reflect phenotypic variation since lost to tend to be fragmentary, (3) there are persistent issues with chronologi- genetic drift and other demographic processes (Lahr, 2016; Lahr & cal dating, and (4) archeological research in the region has a history of Foley, 1998; Manica et al., 2007). However, research has been sty- being underfunded and under-published (Grine, 2016; Lahr, 2016; mied by very small sample sizes with poor geographic and temporal Prendergast & Sawchuk, 2018; Rightmire, 1975). Additionally, only a coverage, and few samples from LSA archeological sites have suffi- small proportion of sites preserve multiple individuals to permit assess- ciently large samples to robustly assess intra-regional population ment of intra-population variability. This is especially true of Late Pleis- variability. tocene and early Holocene skeletons associated with Later Stone Age Here we add Kisese II rockshelter in Tanzania to the list of east- (LSA) foragers that antedate demographic shifts related to the spreads ern Africa sites with LSA skeletal remains. The site preserves a rela- of pastoralism, iron-working, and farming over the past 5000 thousand tively continuous sequence spanning portions of at least the last years (5 ka) as suggested by genetic, skeletal, linguistic, and archeolo- 47 ka that contains faunal remains, stone tools, ochre, and ostrich gical evidence (Marshall & Hildebrand, 2002; Prendergast et al., 2019; eggshell beads (described in Ranhorn et al., n.d. [in review]; Tryon Tishkoff et al., 2009; Wang et al., 2020). Given this paucity of data, et al., 2018). We provide here the first descriptions of six human new samples from LSA archeological contexts are potentially transfor- burials from the site that were excavated in 1956 by Raymond mative for understanding patterns of morphology, biology, and popula- R. Inskeep and present a direct radiocarbon date for one individual tion history among ancient eastern Africans. confirming an early Holocene age and therefore an LSA archeological Eastern African LSA archeological sites with well-described and context. Our objective is to provide the first formal documentation published human remains are limited to fishing localities around Lake and comparison of these individuals, and in doing so contribute to Turkana in northern Kenya (Angel et al., 1980; Barthelme, 1985; Lahr understanding of human morphological variation among past eastern et al., 2016; Phillipson, 1977), Lukenya Hill in Central Kenya African populations. (Gramly, 1976; Gramly & Rightmire, 1973; Tryon et al., 2015), Mumba Rockshelter in northern Tanzania (Bräuer, 1980; Mehlman, 1989), and Mlambalasi Rockshelter in southern Tanzania (Biittner et al., 2017; 2 | M A T E R I A L S A N D M ET H O D S Sawchuk & Willoughby, 2015). Within the Horn of Africa, LSA burials are known from Gogoshiis Qabe in Somalia (Brandt, 1988) and Mota 2.1 | Historical and chronological context in Ethiopia (Arthur et al., 2019; Gallego Llorente et al., 2015). Other LSA burials are found in shell middens around Lake Victoria and are The site of Kisese II is one of several hundred painted rock shelters in attributed to Kansyore fisher-foragers, some of whom may have inter- the Kondoa region of north-central Tanzania (4 290 30.4700 S, acted with herders (Dale & Ashley, 2010; Leakey, 1935; Robertshaw 35 480 43.3100 E; Figure 1), an area with one of the largest concentra- et al., 1983). Depending on one's perception of “eastern” Africa tions of rock art in Africa (Mabulla & Gidna, 2014) and recognized by (e.g., Shea, 2020), this sample could be extended to include individuals UNESCO as the Kondoa Rock Art Sites World Heritage Site. Nearly all from Ishango in the Democratic Republic of the Congo (Crevecoeur of the art at the site is of the naturalistic tradition described by et al., 2016), Gwisho Springs in Zambia (Fagen, 1971), and Hora, Bwasiri and Smith (2015), red in color with human figures and wild Fingira and Chencherere II in Malawi (summarized in Skoglund animals as dominant representations. Initial investigation of the rock et al., 2017). Larger archeological skeletal collections from eastern art in 1935 by L.S.B. and M.D. Leakey led to trial excavations by them Africa are associated with later time periods: early herders living at Kisese II in 1951 (Leakey, 1935, 1983; Leakey & Leakey, 1950). around Lake Turkana
5–4 ka (Hildebrand et al., 2018; Sawchuk Kisese II was targeted because, unusual for the area, it preserved a LAIRD ET AL. 3 F I G U R E 1 Map and Inskeep photos (a) a map of eastern Africa with the location of Kisese II. (b) Kisese II in 2018, and (c) the area approximately corresponding with the area shown in (d–g); photos by Samantha Porter. Photos (d–f) are of Inskeep's excavation of “burial III/IV” and show at least two individuals in spit VII. Ostrich eggshell C14 dates obtained from spit VII suggest two calibrated age ranges that provide minimum age estimates: 12.87–12.15 and 42.79–41.73 ka. The cranium on the right shows fracture patterns and partially-healed cranial trauma that matches juvenile KNM-KX 9 (Figure S3). Notes associated with the skeletal material suggest KNM-KX 1 is also shown in the burial III/IV photo, although the fracture patterns on the ossa coxae visible in the photographs do not precisely match KNM-KX 1. (g) Inskeep's photos show one other burial labeled “burial 2” that contains an older juvenile or adult buried on their side deep (>6-m-thick) stratigraphic sequence that was rich in artifacts and numerals. Isolated human remains in Dar es Salaam were found fossils. Because of this, L.S.B. Leakey hired R.R. Inskeep to conduct during subsequent detailed study of the fauna and have only strati- more extensive excavations there in 1956. graphic information (excavation level) associated with them. Ren- Inskeep is best known for his long and prolific career as an archeologist working in South Africa; Kisese II was his first project as a ewed excavations by Ranhorn at Kisese II in 2017–2019 did not encounter additional burials. field director, and uncharacteristically, is one of the few sites that he excavated without fully publishing (Tryon et al., 2019). Neither the Leakeys nor Inskeep ever published more than a passing reference or 2.2 | Kisese II biometric analyses brief reports on their work at Kisese II, and the presence of burials is not mentioned in any of these publications (Inskeep, 1962; The ages, and in some cases, even stratigraphic contexts of the skele- Leakey, 1935, 1983; Leakey & Leakey, 1950). The only published refer- tal material are largely unknown. Our investigation of the Kisese II ence of the burials is a brief mention by Ambrose (1986), who unsuc- material began in 2011 and includes analyses of archival material from cessfully sampled two of the individuals in the 1980s for isotopic the Leakey and Inskeep excavations. Archival notes found with the analysis. Material from the 1956 excavations was dispersed among specimens suggest Inskeep's spit numbers for some of the burials can multiple institutions (detailed in Tryon et al., 2019). Inskeep notes in an be matched with 14C dates obtained from ostrich eggshell beads with undated draft of a report on Kisese II shared with us by his wife Adi known stratigraphic context (Tryon et al., 2018). Given that the burials (now deceased) that “…the fauna and human remains…were carried by are intrusive and initiated in overlying strata, these dates for the exca- Louis Leakey from Kisese to Nairobi for study and safe keeping.” vation spits provide maximum ages only. The right petrous portions of The human remains were accessioned by the National two individuals (KNM KX 4/5/6 and 7/8) were sampled for ancient Museums of Kenya (NMK) and remain in Nairobi to this day, DNA analysis in 2017; one sample has produced a direct date which although the artifacts and fauna from the 1951 and 1956 excava- we report below. tions were returned to the National Museums of Tanzania (NMT) in As all the burials lack provenience information, individuals were Dar es Salaam in 2012 under the aegis of A. Kwekason. The collec- anatomically associated. Age, sex, and metric analyses for all remains tions in Nairobi are from burials assigned Roman or Arabic were organized by the individual, and the metric comparisons LAIRD ET AL. 4 were focused on the crania and dentition (Tables S1–S6). Skeletal and Hayama, 1982; Leakey, 1935, 1966; Leakey et al., 1943; Leakey fragments from the NMT have not been matched to an individual & Leakey, 1950; Merrick & Monaghan, 1984; Nelson, 1995; (Table S7). Dental wear was scored for the adults using the Phillipson, 1977; Prendergast et al., 2019; Rightmire, 1975; Scott (1979) system that assessed dentine exposure on each molar Robertshaw et al., 1983, 1991; Sassoon, 1968; Sawchuk, 2017; cusp. Dental wear scores were summed to calculate a wear score for Sawchuk & Willoughby, 2015; Sawchuk et al., 2019; Schepartz, 1987; each individual (Table S8). Age was determined for all skeletal material Siiriäinen, 1977; Stiles & Munro-Hay, 1981; Wandibba, 1983; Wang using a combination of dental eruption, suture fusion, and closure of et al., 2020; Table S12). The dental comparative dataset includes an skeletal growth plates (Buikstra & Ubelaker, 1994; Cunningham early pastoralist group representing the first herders in eastern Africa et al., 2016; Ubelaker, 1989; Table S9). Sex was determined for the who lived around Lake Turkana, northern Kenya and pursued a mixed adult material on the ossa coxae using the Phenice (1969) scoring of economy of fishing and herding and possessed novel lithic, ceramic, the ventral arc, subpubic concavity, and ischiopubic ramus ridge, as and well as differences in the greater sciatic notch and preauricular sulcus et al., 2019). Dental metric data were recorded by one author (Buikstra & Ubelaker, 1994). Sex differences in adult skull morphology (Sawchuk) at the National Museums of Kenya and Turkana Basin Insti- were determined by scoring the nuchal crest, mastoid process, supraor- tute (Turkwel facility) in Kenya, the National Museum and House of bital margin, glabella, and mental eminence (Acsadi & Nemeskeri, 1970; Culture in Tanzania, Harvard University's Peabody Museum of Table S10). Sex was not determined for the juvenile remains. Stature Archaeology and Ethnology and the National Museum of Natural estimations for the adults were recorded using measurements from the History in the United States, and the Duckworth Laboratory at the calcaneus and talus (Holland, 1995; Table S11). University of Cambridge in England. The adult dentition from KNM- mortuary traditions (Hildebrand et al., 2018; Sawchuk KX 1, KNM-KX 2, and KNM-KX 4/5/6 were included in the dental comparative analyses (Table S15). 2.3 | Comparative populations Cranial metrics from KNM-KX 2, the only complete undistorted skull, 2.4 | Kisese II craniodental metric analyses were compared to published data from 476 adults from five recent African populations: the San (n = 82) and Zulu (n = 101) from southern A series of 49 measures were recorded from the KNM-KX 1 and KNM- Africa, the Taita (n = 83) from Kenya, a sample from Egypt (n = 111), KX 2 crania, most following Howells (1973, 1989, 1995). Given the and the Dogon (n = 99) from Mali (Howells, 1973, 1989, 1995; https:// fragmentary nature of the Holocene sample, the comparative analyses web.utk.edu/
auerbach/HOWL.htm). The individuals from Howells' focused on nine of the Howells measurements: basion-nasion length dataset were also compared to a sample of 48 African Holocene adults (BNL), biauricular breadth (AUB), bizygomatic breadth (ZYB), glabello- divided into early Holocene LSA (
10.0–4.0 ka; n = 7) and Pastoral occipital length-(GOL), maximum cranial breadth (XCB), minimum cra- Neolithic (
3.5–2.0 ka; n = 41) groups (Angel et al., 1980; Bräuer, nial breadth (WCB), nasal breadth (NLB), nasal height (NLH), and orbit 1983; Gabel, 1965; Leakey, 1935, 1942, 1950; Merrick & Monaghan, height (OBH; Table S14). Unilateral measurements were recorded on 1984; Prendergast et al., 2014; Tables S12 and S13). The early Holo- the left side, but the right side was substituted if measurements on the cene/LSA individuals are geographically, temporally, and culturally left side were not possible. Males and females were analyzed together diverse but are all associated with hunting-fishing-gathering lifeways, to increase the Holocene sample size and because sex determination including ceramic-using Kansyore foragers who may have interacted for KNM-KX 1 and KNM-KX 2 is not definitive. Some measures for the with food producers (Dale & Ashley, 2010). The Pastoral Neolithic comparative individuals represent specialized pastoralists found throughout the (as described in Angel et al., 1980; Bräuer, 1983; Leakey, 1935). sample were approximated due to preservation Central Rift of Kenya and Tanzania who are divided based on mate- Maximum mesiodistal and buccolingual measures were recorded rial culture among other factors into at least two cultural traditions, from all teeth using standard sliding calipers. Third molars were the Elmenteitan and Savanna Pastoral Neolithic, but are genetically excluded from the analyses due to high variability. Measurements were closely related (Prendergast et al., 2019). All measurements for the not recorded for worn teeth (wear ≥ stage 4, per Smith, 1984) or when African Holocene comparative sample were collected from the litera- affected by calculus, debris, or consolidant. Marginally chipped, broken, ture. Early pastoralist (
5.0–4.0 ka) comparative samples were too and reconstructed teeth were recorded when the damage did not fragmented for inclusion in the cranial analyses. Cranial measure- affect the specific measurement taken. Analyses were performed on ments were also recorded from the KNM-KX 1 skull but excluded left teeth, with right antimeres substituted in for missing data wherever from the analyses because of asymmetry (detailed below; Table S14). possible, and males and females were analyzed together. Dental non- Dental metric analyses were undertaken using a sample of metric analyses that include the Kisese II dentition are reported else- 158 African Holocene adults divided into early Holocene/LSA where (Sawchuk, 2017), and adult postcranial measurements are not (
10.0–4.0 ka; n = 37), early pastoralist (
5.0–4.0 ka; n = 32), and analyzed (except for stature) but provided in Table S16. Pastoral Neolithic (
3.5–2.0 ka; n = 89) groups (Ambrose, 1986; Angel All analyses were performed in R (R core team, 2017). Group vari- et al., 1980; Barthelme, 1985; Biittner et al., 2017; Brown, 1966; ance and normality were tested for the cranial and dental metric data Coon, 1971; Hildebrand & Grillo, 2012; Hildebrand et al., 2018; Ikeda using Bartlett's and Shapiro–Wilk's tests. While all measures passed LAIRD ET AL. 5 F I G U R E 2 Cranial remains of KNM-KX 1. The cranium is shown in (a) anterior, (b) posterior, (c) superior, (d) inferior, (e) left lateral, and (f ) right lateral views. The mandible is shown in (g) superior and (h) inferior views. Left (i and j) and right (m and n) zygomatic fragments, with some frontal and maxilla, are shown in anterior and posterior views. Right (k) and left (l) maxillary fragments are shown in lateral view. Right (o) and left (p) scapulae are shown in posterior view. The left (q and r) and right (s and t) ossa coxae are shown in medial and laterial views. The maxillae, scapula, and ossa coxae are not numbered Bartlett's tests (all p > 0.05), some measures were not normally distrib- previously reconstructed using plaster and consolidant resulting in uted and groups' sample sizes were not equal. We used non-parametric artificial cranial deformation and asymmetry. For example, the right Kruskal-Wallis tests to determine whether dental and cranial measure- zygomatic/frontal fragment KNM-KX 3c can be rearticulated at the ments differed between groups. Pairwise comparisons between groups frontal, but the inferior border of the zygomatic touches the petrous were performed using post hoc Dunn tests with a Bonferroni correc- portion of the temporal. This suggests the cranium has been com- tion for multiple comparisons in the R package “dunn.test” (Dinno & pressed anteroposteriorly. The mediolateral reconstruction appears to Dinno, 2017). Significance for all analyses was set at 0.05. be more accurate in that the right and left parietal bosses and changes in the temporal lines occur approximately in the same coronal planes. Mandible KNM-KX 3a is well preserved with full adult dentition miss- 3 | RESULTS ing only the coronoid processes, a small portion of the right gonial angle, the right second incisor and alveolus, and the crown of the right Analyses of the Kisese II rockshelter burials suggest a minimum num- fourth premolar. The mandibular rami have been fragmented and ber of six individuals (Tables S1–S6). The KNM Kisese II collections reconstructed. The mandible occludes with the two maxillary frag- were commingled, and individuals were sorted anatomically. None of ments preserving full adult dentition. Both the mandibular and maxil- the individuals were mineralized, and the physical condition of the lary dentition have similar wear with high dentin exposure (Table S8). specimens were similar to that of the fossil fauna found throughout Several postcranial elements with similar preservation and coloration the stratigraphic sequence. None of the individuals had anatomical are associated with KNM-KX 1. features that fell outside the range of modern African diversity. KNM-KX 2 is well preserved with a complete skull missing only a few teeth (Figure 3; Table S2). The teeth have a lower degree of dental wear compared to KNM-KX 1 (Table S8). The postcrania are also 3.1 | Preservation well preserved although missing elements. The distal left second rib of KNM-KX 2 is enlarged due to osteomyelitis and includes a cloaca KNM-KX 1 consists of a partial skull and postcranial elements (5.82 mm in diameter). A lower right rib also shows pathological (Figure 2; Table S1). Fragments of cranium KNM-KX 1a had been changes associated with osteomyelitis. The shape of the rib deviates LAIRD ET AL. 6 F I G U R E 3 Cranial remains associated with KNM-KX 2. The cranium is shown in (a) anterior, (b) posterior, (c) superior, (d) inferior, (e) left lateral, and (f) right lateral views. The mandible is shown in (g) superior, (h) inferior, (i) left lateral, and (j) right lateral views. The right and left ossa coxae are shown in (k and m) medial and (l and n) lateral views. The mandible and ossa coxae are not numbered to arch medially and the pleural surface of the rib is uneven. Bone brown matrix is present on the cranium and petrous bones and the around this area is smooth indicating that the changes occurred prior cranium has white etching on the left and right parietals. The left fron- to death. This rib also exhibits a series of broad shallow parallel marks tal bone of KNM-KX 9 exhibits partially healed cranial trauma that differ in coloration from the rest of the bone and likely occurred 4.17 mm in diameter (Figure S3C). KNM-KX 10 consists only of maxillae and a mandible preserving postmortem. KNM-KX 4/5/6 had dark brown/orange staining around the some of the dentition (Table S6). The mandible has been fractured in lambdoidal suture and on most of the occipital. Importantly, there are two locations and refit. Individuals KNM-KX 9 and KNM-KX 10 do two associated basiocciputs of similar developmental age indicating not preserve any overlapping elements, and it is possible these ele- that there are at least two individuals associated with KNM-KX 4/5/6 ments belong to the same individual. However, the cranium of KNM- (Table S3). Both basiocciputs lack fusion of the lateral portions (partes KX 9 is larger than KNM-KX 4/5/6, which has a similar age estimate lateralis), and the pars lateralis (shown in Figure S1E and S1F) has as KNM-KX 10 (Table 1). been glued to the pars basilaris. It is possible one of the basiocciputs associated with KNM-KX 4/5/6 belongs to KNM-KX 10, as this individual is also dentally less than 5 years of age (Table 1). Dental wear 3.2 | Chronological dates of the specimens was scored for the adult dentition (Table S8). KNM-KX 7/8 has not undergone reconstruction and some ele- Most of Inskeep's notes associated with the site have been lost, and ments, for example KNM-KX 8a, indicate post-depositional deforma- thus our reconstructions of the age, depth, and archeological associa- tion (Table S4). A gray matrix is adhering to the posterior aspects of tions of the human remains are incomplete. We know the following: the mandible, and a nodule of this matrix is present on the lingual (1) The site was excavated to a depth of
6 m without reaching bed- aspect of the mandibular ramus. The mandible was fractured at the rock. (2) Excavation was done in arbitrary “spits” or levels that were symphysis, and the halves have been glued. assigned Roman numerals from the top downward. These horizontal KNM-KX 9 consists of a partial cranial vault and separate right spits were
15 cm thick, except for the uppermost spits, with spit I and left petrous portions of the temporal bones (Table S5). A dark
59 cm thick and spit II
0.21 cm thick. (3) A series of 29 radiocarbon LAIRD ET AL. TABLE 1 7 Summary of aging methods and estimates for the Kisese II individuals Dental aginga Epiphyseal fusionb Cranial suture closurec Pubic symphysisd Auricular surfacee KNM KX 1/3 21+ 35–43 Phase 4/5: 25–30 Phase 3/4: 30–39 25–43 Phase 3 male-ventral rampart completing, smooth symphysis face, no lipping on margin, and gap on the ventral surface. KNM KX 2 21+ Phase 1:18–19 Phase 2: Age 25–29. Phase 1-female-billowy surface composed of ridges. The upper and lower extremities are not defined. <18 Sphenoccipital synchondrosis: 19–25; Ischial tuberosities: 17–19 Pubic symphysisf Composite age 18–25 KNM-KX 4/5/6 5y − +16mo Basiocciput: <5; Left and right humerus: <11 (male) 3–5 KNM-KX 7/8 Basiocciput: <5; Humerus: <10; Scapula: <14 2y ± 8mo 2y − +8mo KNM-KX 9 KNM-KX 10 <20 <20 3y − +12mo 3y ± 12mo a Ubelaker (1989). Buikstra and Ubelaker (1994). c Meindl and Lovejoy (1985). d Todd (1921a, 1921b). e Lovejoy et al., 1985; Meindl and Lovejoy, 1985; Bedford et al., 1989. f Brooks and Suchey (1990). b dates spanning
4–47 ka on ostrich eggshell fragments from the site Photographs and stratigraphic profiles from Inskeep's archives allow provides a basic chronological framework for the upper 3 m of the us to confidently reconstruct the depth and location only for burial sequence (Tryon et al., 2018). (4) The lower part of the archeological III/IV (found in spit VII). These photographs depict the remains of at sequence can be broadly attributed to the Middle Stone Age, with a least three individuals, of which two were buried on their side in a shift to technologies typical of LSA foragers occurring in spits XVII–XI flexed position (Figure 1). These photographs suggest Inskeep's burial dated to
40–34 ka. While understudied at present, the youngest III/IV corresponds with KNM-KX 9 and possibly KNM-KX 1. Inskeep's spits I and II include a range of ceramic types, diminutive backed photos show skeletal elements (such as a humerus and femur) that microliths of non-local obsidian, and evidence for iron production. were not present in the KNM and NMT Kisese II collections at the Material culture is consistent with occupation of the site by time of study (2014 and 2015). As with much of the non-human fossil populations of foragers who used ceramics (perhaps with Kansyore fauna (Ranhorn et al., n.d. [in review]; Tryon et al., 2019), substantial affinities), pastoralists (broadly attributed to the Pastoral Neolithic), portions of the collection recovered in 1956 are no longer available and iron-equipped agricultural groups beginning in the mid-to-late for study. The finds recovered during faunal analysis that are now in Holocene, broadly within the last 5 ka (Inskeep, 1962; Leakey, 1983; Dar es Salaam do have stratigraphic levels associated with them, and Tryon et al., 2018). The age of the youngest radiocarbon-dated ostrich these suggest that human remains at the site are found as deep as spit eggshell fragment at
4 ka almost certainly does not record the age XIV, some 1.8 m below the 1956 surface of the site. As noted above, of the last use of the shelter. (5) Ochre that may have been used in we assume that all of the burials are intrusive from upper layers, and painting the rock walls is found throughout the excavation, but clear the depths where they were found do not correspond with age, but evidence for the age of the rock art at the site is found only at the rather provide a maximum age for the time of the burial. interface of spits I–II, where a buried painted slab that could be fitted back onto the wall of the shelter was recovered (Inskeep, 1962). Burials III/IV and 7 were each associated with ostrich eggshell beads in the NMK. Ostrich eggshell bead size decreases with time at For the burials, Inskeep was inconsistent in his use of Roman or Kisese II (Tryon et al., 2018), providing a potential relative dating tool Arabic numerals. It is reasonable to assume that they were numbered for these individuals. The beads from burial III/IV are smaller than as they were found, and it may be that burial number correlates with those from burial 7 as expected given the higher stratigraphic position depth, although this is unconfirmed. Handwritten notes in the NMK of burial III/IV. Comparison of bead size suggests that beads with both (presumably from Inskeep) suggest that burial 5 is KNM-KX 10. burials are larger than those above spit VII (Ranhorn et al., n.d. LAIRD ET AL. 8 [in review]). However, as we cannot rule out the possibility that these 5 years ±16 months (Ubelaker, 1989). There are two basiocciputs beads were simply fill from the surrounding sediment rather than associated with this individual, but they both lack fusion of the lateral grave goods associated with the actual burial, a Pleistocene age portions (partes lateralis) suggesting an age of less than five remains an unconfirmed possibility. (Cunningham et al., 2016). No postcranial elements have any epiphy- Given all these uncertainties, direct dates on the Kisese II individ- seal fusion. Of the available elements, Buikstra and Ubelaker (1994) uals are paramount for determining the antiquity of human remains at indicates that humeral distal epiphyseal fusion in males occurs first the site. Due to the delicacy of the collection, only pilot destructive indicating that KNM-KX 4/5/6 is less than age 11. Taken together, sampling was allowed. The petrous portion associated with KNM-KX the cranial and postcranial evidence suggests KNM-KX 4/5/6 is 4/5/6 produced a date of 6210 ± 30 radiocarbon years before pre- between three and 5 years of age. sent (PSUAMS-4718). Following the recommendations of Hogg KNM-KX 7/8: All deciduous teeth were erupted except for the et al. (2020) and using the method reported by Tryon et al. (2018) for canines and second deciduous molars that were partially erupted sub-equatorial tropical regions, we calibrated the radiocarbon date (Figure S2). First adult molar crown was forming in the crypt. This pat- using a mixed model that incorporated data from both the northern tern of dental eruption most closely aligned with individuals aged and southern hemispheres, using OxCal 4.4 software (Ramsey, 2009) 2 years ± 8 months (Ubelaker, 1989). The basiocciput (pars basilaris) and the IntCal20 calibration curves (Hogg et al., 2020; Reimer associated with this individual was unfused from the lateral portions et al., 2020). The resultant estimated age range for the KNM-KX (partes lateralis) suggesting an age of less than five (Cunningham 4/5/6 individual is 7239–6985 cal yr BP at the 95% confidence inter- et al., 2016). The two postcranial elements, a partial right humerus val. This is the first radiocarbon date on human remains from the site and a left scapula, show no epiphyseal fusion suggesting ages of less and establishes an early Holocene/LSA antiquity for at least one of than 10 and 14, respectively. Individual KNM-KX 7/8 is likely aged the burials and potentially others. 2 years ± 8 months. KNM-KX 9: Age was not easily assessed for KNM-KX 9 because of the relative paucity of materials (Figure S3). The coronal suture 3.3 | Age does not show any age-related closure suggesting the individual was 20 years old or younger (Meindl & Lovejoy, 1985). Specimen KNM- KNM-KX 1: The mandible and maxilla preserve fully erupted adult KX 9a/8c preserves the left portion of the sagittal suture posterior to dentition. Age was assessed using the closure and fusion of cranial bregma but the degree of closure cannot be assessed. Age was not sutures indicating an age range of 35–43 years (Buikstra & estimated from the petrous fragments. Ubelaker, 1994; Meindl & Lovejoy, 1985). The pubic symphysis cor- KNM-KX 10: All visible deciduous mandibular and maxillary denti- responded with Phase 4/5 and an age range of 25–30 years. Similarly, tion were fully erupted and the crown of the first adult molars are visi- the auricular surface was estimated to be Phase 3/4 and an age range ble in their crypts (Figure S4). Using Ubelaker (1989), age of KNM-KX of 30–39. KNM-KX 1 has an estimated age range of 25–43 years. 10 was estimated as 3 years ±12 months. KNM-KX 2: Cranial and postcranial remains of KNM-KX 2 are well preserved. Adult mandibular and maxillary dentition are fully erupted with occlusal wear. The spheno-occipital synchondrosis is partially fused indi- 3.4 | Sex cating an age of 18–24 or potentially younger (Cunningham et al., 2016). Age was also assessed using cranial suture closure (following Buikstra & Sex was determined for the two adult individuals, KNM-KX 1 and Ubelaker, 1994; Meindl & Lovejoy, 1985), and KNM-KX 2a was assessed KNM-KX 2, from the skull and os coxae (Table 2). We did not assess to have no suture closure corresponding with an age of less than 18 (- sex for the juvenile individuals. Table S9). Age was assessed from right os coxae using three methods. KNM-KX 1: Sex was estimated from the skull using scoring Secondary ossification centers on the iliac crest and the ischial tuberosity detailed in Acsadi and Nemeskeri (1970). Scoring of the nuchal crest are partially fused indicating an age range from 14 to 22 years (Buikstra & and glabella was ambiguous, and the mastoid process and mental emi- Ubelaker, 1994). An age range of 18–19 was estimated from the pubic nence were possibly female. The supraorbital margin was not scored. symphysis of the left os coxae (Table 1; Brooks & Suchey, 1990; Katz & Sex for the skull was indeterminate. On the os coxae, the ventral arc, Suchey, 1986; Todd, 1921a, 1921b). The auricular surface corresponded subpubic concavity and medial ridge were not preserved. The greater with Phase 1–2 and an age estimate ranging from 20 to 29 years (Bedford sciatic notch was scored on the right os coxae fragment as narrow indi- et al., 1989; Lovejoy et al., 1985). Collectively, age estimates for KNM-KX cating male, and a preauricular sulcus was absent suggesting the indi- 2 range from 18 to 29 years but a younger range of 18–25 is more likely vidual is male (Karsten, 2018). Individual KNM-KX 1 is possibly male. based on the lack of fusion of the spheno-occipital synchondrosis. KNM-KX 2: Sex determination from the skull followed Acsadi and KNM-KX 4/5/6: The mandible (KNM-KX 6a/6b) preserves the Nemeskeri (1970). The nuchal crest, supraorbital margin, and mental fully occluded and worn deciduous first and second molars as well as eminence were ambiguous, but the left mastoid process and glabella the crown of the adult right canine, the adult left first incisor, and a trended toward female. Sex from the skull is possibly female. The ven- crypt for the left adult second molar (Figure S1). Based on the avail- tral arc, subpubic concavity and the medial ridge were scored on the able dentition, this eruption pattern is most similar to individuals aged left os coxae as possibly males (Table 2). The greater sciatic notch was LAIRD ET AL. 9 Summary of methods and estimates of sex for the Kisese II individuals TABLE 2 Skull morphologya Subpubic regionb Greater sciatic notchc Preauricular sulcusd Sex determination KNM KX 1/3 Indeterminate - Narrow notch: 5; M Absent: 0, M Possibly male KNM KX 2 F? Ventral arc: 2; Subpubic concavity: 3; Ischiopubic ramus: 3; M? Wide notch: 2; F? Wide sulcus: 2 Possibly female a Acsadi and Nemeskeri (1970). Phenice (1969). c Walker (2005). d Karsten (2018). b wide indicating female, and a wide preauricular sulcus was present. Holocene/LSA (
10.0–4.0 ka) individuals had the largest teeth Individual KNM-KX 2 is possibly female. followed by early pastoralists. The smallest teeth are found in the Pastoral Neolithic (
3.5–2.0 ka) sample (Figure 4). With the exception of the mesiodistal length of the upper canine and lower second incisor, 3.5 | Stature there were no significant differences in measurements for the incisors and canines. Dental measures varied significantly for upper third and Despite their presence in photos from the excavation, no adult long fourth premolar mesiodistal length and the upper fourth premolar bones were found in the Kisese II collections at NMK or NMT. Stat- buccolingual width (Kruskal-Wallis: χ2 = 16.00–9.52, df = 3, ure estimates for the adults were based on measurements of the p < 0.02). Only buccolingual width of the lower third premolar varied calcaneus and talus for African-American females and males significantly in the lower premolars (χ2 = 10.00, df = 3, p = 0.02). (Holland, 1995). These measures suggest an average stature of However, pairwise group comparisons for the premolars indicated 167.86 (±5.38) cm, a minimum stature of 154.45 cm, and a maxi- the Kisese II samples did not significantly differ from the other mum stature of 172.28 cm for the individual labeled KNM-KX groups (all p > 0.05). 1 (Table S11). The burial labeled KNM-KX 2 had an average stature Buccolingual and mesiodistal measures from the upper and lower of 167.79 (±5.57), a minimum stature of 154.00 cm, and a maximum first and second molars significantly differed across groups of 173.22 cm (Table S11). (χ2 = 11.52–24.48, df = 3, p < 0.01), with the exception of mesiodistal length of the upper first molar (χ2 = 1.37, df = 3, p = 0.71). The Kisese II upper first molar buccolingual width was significantly smaller than 3.6 | Comparative cranial metric analyses the early Holocene/LSA (p = 0.02) and early pastoralist (p = 0.01) samples but did not differ from the Pastoral Neolithic group (p = 0.42). Cranial measurements from KNM-KX 2 were compared to early Pairwise comparisons suggest the Kisese II dentitions did not differ Holocene/LSA (
10.0–4.0 ka), Pastoral Neolithic (
3.5–2.0 ka), from the other groups for the upper second molar buccolingual and and modern African groups (Figure 4). The nine cranial measure- mesiodistal measurements (both p > 0.05). Pairwise comparisons indi- ments included in the comparative sample varied significantly cate the Kisese II buccolingual measurements for the first and second across the population groups (Kruskal-Wallis: χ2 = 51.96–202.27, lower molars were significantly smaller than the early Holocene/LSA df = 7, p < 0.01). Pairwise tests were not used for the cranial metric group (LM1 p = 0.05; LM2 p = 0.03). analyses because the Kisese II sample only consisted of one individual. The Taita, Early Holocene/LSA, Pastoral Neolithic, and KNMKX 2 all had a similar ratio of maximum cranial breadth and length 4 | DI SCU SSION AND CO NCLUSIO NS compared to the other modern African populations. Dimensions of the nasal aperture for KNM-KX 2 were smaller than most of the Kisese II rockshelter has yielded artifacts and faunal remains that span modern African populations but overlapped with Egyptian individ- much of the Late Pleistocene and Holocene and document a number uals. Kruskal-Wallis tests for all cranial measurements across indi- of archeological transitions in eastern Africa (Ranhorn et al., n.d. [in vidual sites within the early Holocene/LSA and Pastoral Neolithic review]; Tryon et al., 2018). Here we provide the first descriptions of samples were not significant (all p > 0.05). the human remains found at the site, which represent at least six individuals encompassing both sexes and ranging in age from young children to middle-aged adults. It is important to stress that we have 3.7 | Comparative dental metric analyses adopted as an initial approach one that emphasizes morphology and comparison at the population level rather than one that seeks to Although there was overlap between groups, most mesiodistal and understand the lived experiences of those buried at Kisese II. Given buccolingual measures varied chronologically such that the early the history and nature of this legacy skeletal collection, our first goal LAIRD ET AL. 10 F I G U R E 4 Cranial and dental metric comparisons. Boxplots showing mandibular (a) and maxillary (b) buccolingual widths. Boxplots of the ratio of maximum cranial breadth to length (c) and nasal breadth to length (d). Boxplots showing the ratios maximum cranial breadth to length (e) and nasal breadth to length (f) in the early Holocene later stone age (LSA) and pastoral Neolithic samples. For all plots, the upper and lower bound of the boxes corresponds with the 25th and 75th percentiles and the whiskers extend 1.5 times the interquartile range in either direction. The median is represented by a horizontal line inside the boxes is to describe individuals excavated more than 50 years ago who have calculus and ancient DNA) may well provide a more nuanced under- largely been forgotten since. Future archival work, stratigraphic and standing of these early occupants of the Kondoa region. contextual information from renewed excavations at Kisese II, and Our results indicate that individuals buried at Kisese II signifi- other approaches (e.g., isotopic analyses and/or studies of dental cantly expand the range of morphological variation of eastern African LAIRD ET AL. 11 human populations during the Holocene and perhaps Late Pleisto- the presence of only one comparable adult cranium, KNM-KX 2 was cene. Our interpretations are necessarily tempered by two limitations: within the range of variation of modern Africans and overlapped with (1) With the exception of KNM-KX 4/5/6 at
7.1 ka, the Kisese II the early Holocene and Pastoral Neolithic comparative samples. Of individuals as yet lack firm relative or chronometric dates and cannot note, KNM-KX 2 differed from the burials at the nearby site of be demonstrably associated with particular artifact types. (2) Photos Mumba, particularly in nasal dimensions. These results suggest that from Inskeep's excavations show that adult long bones were present KNM-KX at the time of excavation, but these elements have been subsequently geographicallysimilar sites. These analyses highlight variation in Late lost. The collection available for study is therefore only a portion of Pleistocene-early Holocene humans and may indicate that LSA east- what was recovered in 1956. This is, unfortunately, a pattern charac- ern African foragers possibly exhibited greater morphological variation teristic of much of the Kisese II assemblage as a whole (detailed in relative to contemporary Africans. These results mirror high levels of Tryon et al., 2019). material cultural variation observed among LSA sites and potentially 2 differs morphologically from chronologically and These same archival photographs do, however, indicate that at support interpretations of increasing regionalization seen in the arche- least two individuals were buried on their side in a flexed position. ological record (for discussion, see Tryon & Faith, 2013; Tryon & Flexed burial postures date to
15 ka in Tanzania with the oldest evi- Ranhorn, 2020; Wilshaw, 2016). dence from Mlambalasi in the Iringa region
350 km to the south Individuals buried at Kisese II provide an opportunity to explore (Sawchuk & Willoughby, 2015), and are noted at the closer (
80 km) human regional variation during the Holocene and potentially the Late site of Mumba (Bräuer, 1980). Ostrich eggshell beads were also pre- Pleistocene. Collectively, our metric analyses suggest (1) substantial sent in levels associated with the burials at all three sites (Biittner variation in craniodental morphology among Holocene and possibly et al., 2017; Bushozi, 2020; Mehlman, 1979; Tryon et al., 2018). Evi- Late Pleistocene eastern African populations, but also (2) morphologi- dence from Kisese II may reflect a shared mortuary behavior in the cal similarity at localized sites like Kisese II during the Holocene. Con- region. tinued exploration of Kisese II's archival and excavated materials will Craniodental comparisons highlight similarities between the improve not only our understanding of the burials' archeological con- Kisese II sample and eastern African Pastoral Neolithic groups. The text at the site, but also regional variation in human morphology and Kisese II deposits suggest discontinuous human occupation over at behavior in eastern Africa among LSA sites. Future research that com- least the last 47 ka, occupied by populations using stone, ceramic, bines morphological data with other lines of evidence, such as regional and iron technologies that hunted but who used domestic stock differences in how stone or ceramic artifacts were made and used (e.g., sheep or goat) at least occasionally (Tryon, 2019). We cannot (Ashley & Grillo, 2015; Seitsonen et al., 2013; Wilshaw, 2016), pat- associate individual burials with particular technologies or archeolo- terns of long distance raw material exchange (e.g., Frahm & gical entities, but comparison of the date of
7.1 ka for KNM-KX 4/ Tryon, 2018; Mehlman, 1989; Prendergast et al., 2013), burial prac- 5/6 with established regional chronologies firmly indicates that this tices, and aDNA recovery (Prendergast et al., 2019; Wang individual antedates the introduction of either domestic stock or iron et al., 2020), has immense potential for reconstructing complex social into the region, and can instead be associated with either aceramic interactions during this time in the African past. or potentially ceramic (cf. Kansyore) LSA foragers (reviewed in Mehlman, 1979; Prendergast et al., 2014). With a date of
7.1 ka, ACKNOWLEDG MENTS dental measurements of KNM-KX 4/5/6 are expected to be most This project was supported by funding through Harvard University, similar to the early Holocene comparative sample. However, the American School for Prehistoric Research, the Leakey Founda- buccolingual and mesiodistal dental measures of KNM-KX 4/5/6, as tion, the New York University Cliff Jolly award, the Rutgers Univer- well as KNM-KX 1 and KNM-KX 2, were closest to the Pastoral Neo- sity Byrne Seminar Research Program, NSF IGERT DGE 0801634, lithic sample, broadly dated from 4–1.5 ka (Figure 4). This suggests Fulbright-Hays DDRA, and the Social Sciences and Humanities the individuals from Kisese II had relatively smaller dentitions than Research Council of Canada (grant 767-2012-1903). Analyses of the early Holocene foragers, but similar to those of early pastoralist and Kisese II skeletal material were undertaken with permission from the Pastoral Neolithic eastern Africans. If KNM-KX 1 and KNM-KX 2 are Tanzania Commission for Science and Technology (COSTECH) under substantially younger than KNM-KX 4/5/6, the Kisese II samples permit numbers: 2014-233-NA-2013-122; 2015-115-ER-2013-122, would imply the relative persistence of small teeth at the site across 2015-116-ER-2015-212, 2015-120-NA-2015-24, and permit num- the Holocene. ber NCST/5/002/R/576 issued by the Kenyan National Commission Our findings contribute to a growing body of evidence that sug- for Science, Technology, and Innovation (NACOSTI). Dental metric gests that human skeletal remains from the Late Pleistocene and early data were collected under NACOSTI (NACOSTI/P/14/1876/1410) Holocene in eastern Africa exhibit relatively high amounts of morpho- and COSTECH (2013-223-NA-2014-101) permits. These results logical diversity (Crevecoeur et al., 2009; Crevecoeur et al., 2016; have been presented to stakeholders living near the Kisese II site Grine et al., 2007; Mounier et al., 2018; Stojanowski, 2014). Com- (the Machinjioni village council) and are published with their permis- pared to other eastern Africans, KNM-KX 2 had a similar cranial sions. Thanks to the staff of the National Museum of Tanzania, the breadth and length, but relatively small dentition, and a relatively large National Museums of Kenya, and Fredrick Manthi for facilitating this nasal aperture. While our analyses of cranial metrics were limited by study. We thank David Reich, Mary Prendergast, Douglas Kennett, LAIRD ET AL. 12 and Brendan Culleton for the radiocarbon date, and acknowledge support from an NSF Archaeometry program grant to Kennett and Culleton (BCS-1460369). Thanks to Samantha Porter for providing photos from the 2018 Kisese II excavations, and Jennifer Charlson for permission to publish photographs from the Inskeep archives. Finally, thanks to Tyler Faith for discussion, and Isabelle Crevecoeur and Hugo Reyes-Centero for their assistance with the comparative material. AUTHOR CONTRIBUTIONS Myra F. Laird: Conceptualization; data curation; formal analysis; investigation; methodology; writing-original draft; writing-review & editing. Elizabeth Sawchuk: Data curation; funding acquisition; investigation; resources; writing-original draft; writing-review & editing. Amandus Kwekason: Project administration; resources; writing-review & editing. Audax Mabulla: Project administration; resources; writingreview & editing. Emmanuel Ndiema: Resources; writing-review & editing. Christian Tryon: Conceptualization; funding acquisition; investigation; project administration; resources; writing-original draft; writing-review & editing. 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